Biology:Chlorophyta

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Short description: Phylum of green algae

Chlorophyta
EB1911 Algae - Fig. 2.-Chlorophyceae.png
Chlorophytes (A–F, H–L and O)
Scientific classification e
Domain: Eukaryota
Clade: Diaphoretickes
(unranked): Archaeplastida
(unranked): Viridiplantae
Division: Chlorophyta
Reichenbach, 1828, emend. Pascher, 1914, emend. Lewis & McCourt, 2004[1][2][3]
Classes[4]
Synonyms[5]
  • Chlorophycophyta Papenfuss 1946
  • Chlorophycota
  • Chlorophytina
  • Chlorophyllophyceae
  • Isokontae
  • Stephanokontae
Green algae on coastal rocks at Shihtiping in Taiwan

Chlorophyta is a taxon of green algae informally called chlorophytes.[6] The name is used in two very different senses, so care is needed to determine the use by a particular author. In older classification systems, it is a highly paraphyletic group of all the green algae within the green plants (Viridiplantae) and thus includes about 7,000 species[7][8] of mostly aquatic photosynthetic eukaryotic organisms. In newer classifications, it is the sister clade of the streptophytes/charophytes. The clade Streptophyta consists of the Charophyta in which the Embryophyta (land plants) emerged.[9][10] In this latter sense the Chlorophyta includes only about 4,300 species.[4] About 90% of all known species live in freshwater.[11] Like the land plants (embryophytes: bryophytes and tracheophytes), green algae (chlorophytes and charophytes besides embryophytes) contain chlorophyll a and chlorophyll b and store food as starch[7] in their plastids.

With the exception of the three classes Ulvophyceae, Trebouxiophyceae and Chlorophyceae in the UTC clade, which show various degrees of multicellularity, all the Chlorophyta lineages are unicellular.[12] Some members of the group form symbiotic relationships with protozoa, sponges, and cnidarians. Others form symbiotic relationships with fungi to form lichens, but the majority of species are free-living. Some conduct sexual reproduction, which is oogamous or isogamous. All members of the clade have motile flagellated swimming cells.[13] While most species live in freshwater habitats and a large number in marine habitats, other species are adapted to a wide range of land environments. For example, Chlamydomonas nivalis, which causes Watermelon snow, lives on summer alpine snowfields. Others, such as Trentepohlia species, live attached to rocks or woody parts of trees. Monostroma kuroshiense, an edible green alga cultivated worldwide and most expensive among green algae, belongs to this group.

Ecology

Species of Chlorophyta (treated as what is now considered one of the two main clades of Viridiplantae) are common inhabitants of marine, freshwater and terrestrial environments.[14][15] Several species have adapted to specialised and extreme environments, such as deserts, arctic environments, hypersaline habitats, marine deep waters, deep-sea hydrothermal vents and habitats that experiences extreme changes in temperature, light and salinity.[16][17][18][19] Some groups, such as the Trentepohliales are exclusively found on land.[20] Several species of Chlorophyta live in symbiosis with a diverse range of eukaryotes, including fungi (to form lichens), ciliates, forams, cnidarians and molluscs. [15] Some species of Chlorophyta are heterotrophic, either free-living or parasitic.[21][22] Others are mixotrophic bacterivores through phagocytosis.[23] Two common species of the heterotrophic green alga Prototheca are pathogenic and can cause the disease protothecosis in humans and animals.[24]

Classifications

"Siphoneae" from Ernst Haeckel's Kunstformen der Natur, 1904

Characteristics used for the classification of Chlorophyta are: type of zoid, mitosis (karyokinesis), cytokinesis, organization level, life cycle, type of gametes, cell wall polysaccharides[25] and more recently genetic data.

Phylogeny

Leliaert et al. 2012 proposed the following phylogeny. He marked the "prasinophytes" as paraphyletic, with the remaining Chlorophyta groups as "core chlorophytes". He described all Streptophyta except the land plants as paraphyletic "charophytes".[15]

"Hypothetical ancestral
 
Chlorophyta
 
 
 

Palmophyllales

 
 

Prasinococcales

 
 

Nephroselmidophyceae

 
 
 
 
 

Pycnococcaceae

 
 
 

Pyramimonadophyceae

 
 

Mamiellophyceae

 
 
 
 
 

Picocystis

 
 
 

Pedinophyceae

 
 
 

Chlorodendrophyceae

 
 
 

Trebouxiophyceae

 
 
 

Ulvophyceae

 
 

Chlorophyceae

 
 
 
 
 
 
 
 
 
Streptophyta
 
 

Mesostigmatophyceae

 
 

Chlorokybophyceae

 
 
 
 

Klebsormidiophyceae

 
 
 

Charophyceae (stoneworts)

 
 
 
 

Zygnematophyceae

 
 

Coleochaetophyceae

 
 
 

Embryophyte (Land plants)

 
 
 
 
 
 
Script error: No such module "Clade/labels".
green flagellate"

A 2020 paper places the "Prasinodermophyta" (i.e. Prasinodermophyceae + Palmophyllophyceae) as the basal Viridiplantae clade.[26]

Viridiplantae/
 
Prasinodermophyta
 

Prasinodermophyceae

 
Palmophyllophyceae
 

Prasinococcales ("prasinophyte clade VI")

 
 

Palmophyllales

 
 
 
 
 
Chlorophyta
 
 

Pyramimonadophyceae ("prasinophyte clade I")

 
 

Mamiellophyceae ("prasinophyte clade II")

 
 
 
 

Nephroselmidophyceae ("prasinophyte clade III")

 
 
 

Pycnococcaceae ("prasinophyte clade V")

 
 
 

Chloropicophyceae ("prasinophyte clade VII A/B/C")

 
Tetraphytina
 

Pedinophyceae

 
 
 

Trebouxiophyceae

 
 
 

Chlorodendrophyceae ("prasinophyte clade IV")

 
 
 

Ulvophyceae

 
 

Chlorophyceae

 
 
 
 
 
 
 
 
 
Streptophyta/
 
Mesostigmatophyceae
 

Mesostigma viride

 
 
 

Spirotaenia

 
 

Chlorokybophyceae

 
 
 
 
 
 

Klebsormidiophyta

 
Phragmoplastophyta
 

Charophyceae (stoneworts)

 
 
Coleochaetophyceae
 

Chaetosphaeridiales

 
 

Coleochaetophyta

 
 
 
Zygnematophyceae
 

Zygnematophyta

 
 

Mesotaeniaceae

 
 
 

Embryophyte (Land plants)

 
 
 
 
 
Charophyta
 
Green Algae/
Prasinophyta s.l.

Leliaert et al. 2012

Simplified phylogeny of the Chlorophyta, according to Leliaert et al. 2012.[15] Note that many algae previously classified in Chlorophyta are placed here in Streptophyta.

Pombert et al. 2005

A possible classification when Chlorophyta refers to one of the two clades of the Viridiplantae is shown below.[27]

Lewis & McCourt 2004

Hoek, Mann and Jahns 1995

Classification of the Chlorophyta, treated as all green algae, according to Hoek, Mann and Jahns 1995.[7]

In a note added in proof, an alternative classification is presented for the algae of the class Chlorophyceae:

Bold and Wynne 1985

Classification of the Chlorophyta and Charophyta according to Bold and Wynne 1985.[29]

Mattox & Stewart 1984

Classification of the Chlorophyta according to Mattox & Stewart 1984:[28]

Fott 1971

Classification of the Chlorophyta according to Fott 1971.[7]:483

  • Suborder Chlorosarcineae
  • Suborder Ulotrichineae
  • Suborder Oedogoniineae
  • Suborder Chaetophorineae

Round 1971

Classification of the Chlorophyta and related algae according to Round 1971.[30]

Smith 1938

Classification of the Chlorophyta according to Smith 1938:

  • Order 1. Volvocales
  • Order 5. Schizogoniales
  • Family Schizogoniaceae
  • Order 10. Siphonales
  • Order 11. Siphonocladiales

Research and discoveries

In February 2020, the fossilized remains of green algae, named Proterocladus antiquus were discovered in the northern province of Liaoning, China.[31] At around a billion years old, it is believed to be one of the oldest examples of a multicellular chlorophyte.[32]

References

  1. ↑ Conspectus Regni Vegetabilis. 1828. p. 23. https://books.google.com/books?id=OWkVAAAAYAAJ&pg=PA23. 
  2. ↑ "Über Flagellaten und Algen". Berichte der Deutschen Botanischen Gesellschaft 32: 136–160. 1914. doi:10.1111/j.1438-8677.1914.tb07573.x. https://www.biodiversitylibrary.org/item/138036#page/145/mode/1up. 
  3. ↑ "The new higher level classification of eukaryotes with emphasis on the taxonomy of protists". The Journal of Eukaryotic Microbiology 52 (5): 399–451. 2005. doi:10.1111/j.1550-7408.2005.00053.x. PMID 16248873. 
  4. ↑ 4.0 4.1 "AlgaeBase : Chlorophyta". World-wide electronic publication, National University of Ireland, Galway. 2011. http://www.algaebase.org/browse/taxonomy/?searching=true&gettaxon=Chlorophyta. 
  5. ↑ "The Classification of the Algae". A century of progress in the natural sciences, 1853-1953. California Academy of Sciences. 1955. https://archive.org/stream/centuryofprogres00cali#page/126/mode/1up. 
  6. ↑ "The phycocyanobilin chromophore of streptophyte algal phytochromes is synthesized by HY2". The New Phytologist 214 (3): 1145–1157. May 2017. doi:10.1111/nph.14422. PMID 28106912. 
  7. ↑ 7.0 7.1 7.2 7.3 Algae An Introduction to Phycology. Cambridge: Cambridge University Press. 1995. ISBN 978-0-521-30419-1. https://books.google.com/books?id=xuUoiFesSHMC. 
  8. ↑ "Major Algae Phyla - Table - MSN Encarta". http://encarta.msn.com/media_461543986/Major_Algae_Phyla.html. 
  9. ↑ "Green algae and the origin of land plants". American Journal of Botany 91 (10): 1535–56. October 2004. doi:10.3732/ajb.91.10.1535. PMID 21652308. 
  10. ↑ "Streptophyte algae and the origin of embryophytes". Annals of Botany 103 (7): 999–1004. May 2009. doi:10.1093/aob/mcp044. PMID 19273476. 
  11. ↑ Lee, Robert Edward (2018). Chlorophyta (Chapter 5) - Phycology - Cambridge University Press. doi:10.1017/9781316407219. ISBN 9781316407219. https://www.cambridge.org/core/books/phycology/chlorophyta/5D438D2AA6C763806B876D804C511A0E. 
  12. ↑ "Green algae and the origins of multicellularity in the plant kingdom". Cold Spring Harbor Perspectives in Biology 6 (11): a016170. October 2014. doi:10.1101/cshperspect.a016170. PMID 25324214. 
  13. ↑ "Nuclear DNA content estimates in green algal lineages: chlorophyta and streptophyta". Annals of Botany 99 (4): 677–701. April 2007. doi:10.1093/aob/mcl294. PMID 17272304. 
  14. ↑ Graham LE, Graham JM, Wilcox LW (2009) Algae. 2nd Edition. Benjamin Cummings (Pearson), San Francisco, CA
  15. ↑ 15.0 15.1 15.2 15.3 "Phylogeny and molecular evolution of the green algae". Critical Reviews in Plant Sciences 31: 1–46. 2012. doi:10.1080/07352689.2011.615705. http://images.algaebase.org/pdf/5628E58F0ecc431F0CsJm2B04CAD/49951.pdf. 
  16. ↑ "Unearthing the molecular phylodiversity of desert soil green algae (Chlorophyta)". Systematic Biology 54 (6): 936–47. December 2005. doi:10.1080/10635150500354852. PMID 16338765. 
  17. ↑ "Hidden levels of phylodiversity in Antarctic green algae: further evidence for the existence of glacial refugia". Proceedings. Biological Sciences 276 (1673): 3591–9. October 2009. doi:10.1098/rspb.2009.0994. PMID 19625320. 
  18. ↑ "Into the deep: new discoveries at the base of the green plant phylogeny". BioEssays 33 (9): 683–92. September 2011. doi:10.1002/bies.201100035. PMID 21744372. 
  19. ↑ "The unexpected extremophile: Tolerance to fluctuating salinity in the green alga Picochlorum". Algal Research 16: 465–472. June 2016. doi:10.1016/j.algal.2016.04.003. 
  20. ↑ "Molecular systematics of the subaerial green algal order Trentepohliales: an assessment based on morphological and molecular data". International Journal of Systematic and Evolutionary Microbiology 56 (Pt 7): 1709–15. July 2006. doi:10.1099/ijs.0.63990-0. PMID 16825655. 
  21. ↑ "Parasitic green algae". Annu. Rev. Phytopathol. 9: 45–64. 1971. doi:10.1146/annurev.py.09.090171.000401. 
  22. ↑ "Complex patterns of plastid 16S rRNA gene evolution in nonphotosynthetic green algae". Journal of Molecular Evolution 53 (6): 670–9. December 2001. doi:10.1007/s002390010254. PMID 11677627. Bibcode: 2001JMolE..53..670N. 
  23. ↑ Anderson, R.; Charvet, S.; Hansen, P. J. (2018). "Mixotrophy in Chlorophytes and Haptophytes—Effect of Irradiance, Macronutrient, Micronutrient and Vitamin Limitation - NCBI - NIH". Frontiers in Microbiology 9: 1704. doi:10.3389/fmicb.2018.01704. PMID 30108563. 
  24. ↑ "Phylogenetic analysis identifies the invertebrate pathogen Helicosporidium sp. as a green alga (Chlorophyta)". International Journal of Systematic and Evolutionary Microbiology 52 (Pt 1): 273–9. January 2002. doi:10.1099/00207713-52-1-273. PMID 11837312. 
  25. ↑ Lobban, Christopher S.; Wynne, Michael James (1981). The Biology of Seaweeds.. Botanical Monograph Series 17.. University of California Press. p. 88. ISBN 9780520045859. 
  26. ↑ "The genome of Prasinoderma coloniale unveils the existence of a third phylum within green plants". Nature Ecology & Evolution 4 (9): 1220–1231. September 2020. doi:10.1038/s41559-020-1221-7. PMID 32572216. 
  27. ↑ "The chloroplast genome sequence of the green alga Pseudendoclonium akinetum (Ulvophyceae) reveals unusual structural features and new insights into the branching order of chlorophyte lineages". Molecular Biology and Evolution 22 (9): 1903–18. September 2005. doi:10.1093/molbev/msi182. PMID 15930151. 
  28. ↑ 28.0 28.1 28.2 28.3 28.4 "Classification of the green algae: a concept based on comparative cytology.". The systematics of Green Algae. 27. London: Academic Press. 1984. pp. 29–72. 
  29. ↑ Introduction to the algae : structure and reproduction (2nd ed.). Englewood Cliffs, N.J.: Prentice-Hall. 1985. ISBN 978-0-13-477746-7. 
  30. ↑ "The taxonomy of the Chlorophyta, 2". Br. Phycol. J. 6 (2): 235–264. 1971. doi:10.1080/00071617100650261. 
  31. ↑ "Billion-year-old green algae found in China is the oldest ever discovered" (in en). 2020-02-24. https://www.newsweek.com/billion-year-old-green-algae-china-oldest-1488794. 
  32. ↑ Sandhya Ramesh (2020-02-25). "Scientists discover world's oldest green algae fossil in China" (in en-US). https://theprint.in/science/scientists-discover-worlds-oldest-green-algae-fossil-in-china/370404/. 

Further reading

  • Seaweeds of the British Isles. 2 (Chlorophyta). London: Natural History Museum. 1991. ISBN 978-0-565-00981-6. 
  • "Green algae and the origin of land plants". American Journal of Botany 91 (10): 1535–56. October 2004. doi:10.3732/ajb.91.10.1535. PMID 21652308. 
  • Green Algae. Structure, Reproduction and Evolution in Selected Genera. Stamford, CT: Sinauer Assoc.. 1975. pp. 606. 

Wikidata ☰ Q264543 entry



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